CINXE.COM

<!DOCTYPE html><html lang="en-US"><head><title>Just a moment...</title><meta http-equiv="Content-Type" content="text/html; charset=UTF-8"><meta http-equiv="X-UA-Compatible" content="IE=Edge"><meta name="robots" content="noindex,nofollow"><meta name="viewport" content="width=device-width,initial-scale=1"><style>*{box-sizing:border-box;margin:0;padding:0}html{line-height:1.15;-webkit-text-size-adjust:100%;color:#313131;font-family:system-ui,-apple-system,BlinkMacSystemFont,Segoe UI,Roboto,Helvetica Neue,Arial,Noto Sans,sans-serif,Apple Color Emoji,Segoe UI Emoji,Segoe UI Symbol,Noto Color Emoji}body{display:flex;flex-direction:column;height:100vh;min-height:100vh}.main-content{margin:8rem auto;max-width:60rem;padding-left:1.5rem}@media (width <= 720px){.main-content{margin-top:4rem}}.h2{font-size:1.5rem;font-weight:500;line-height:2.25rem}@media (width <= 720px){.h2{font-size:1.25rem;line-height:1.5rem}}#challenge-error-text{background-image:url(data:image/svg+xml;base64,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);background-repeat:no-repeat;background-size:contain;padding-left:34px}@media (prefers-color-scheme:dark){body{background-color:#222;color:#d9d9d9}}</style><meta http-equiv="refresh" content="390"></head><body class="no-js"><div class="main-wrapper" role="main"><div class="main-content"><noscript><div class="h2"><span id="challenge-error-text">Enable JavaScript and cookies to continue</span></div></noscript></div></div><script>(function(){window._cf_chl_opt={cvId: '3',cZone: "onlinelibrary.wiley.com",cType: 'managed',cRay: '8fd943f7bfa9021f',cH: 'zB8uGckB4m2YtC4V6dDO7.1um2V2ueNoJL_oK4KKj80-1736140895-1.2.1.1-fSw5s5tWGTKtGYC1HTcG4b7z6MU5.ZufgXpJXRT.t6xErtoVSxb81xt4t8Ye1J7V',cUPMDTk: "\/doi\/abs\/10.1111\/gcb.15913?__cf_chl_tk=vIEGH1hh8sID48KObbzsNggu0n2iA_nbvt72Y_p.bHk-1736140895-1.0.1.1-2JAYcK59dTpFbzC1mwZe35ArUNa7OBY1gZaNtF2oSSs",cFPWv: 'g',cITimeS: '1736140895',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "",fa: "\/doi\/abs\/10.1111\/gcb.15913?__cf_chl_f_tk=vIEGH1hh8sID48KObbzsNggu0n2iA_nbvt72Y_p.bHk-1736140895-1.0.1.1-2JAYcK59dTpFbzC1mwZe35ArUNa7OBY1gZaNtF2oSSs",md: "pjcEFRNCBn6PuJAQGzf3RxVhbxOvVhdUanehDAIvdeU-1736140895-1.2.1.1-P0r6ZlLhvOzdth1hsh9QM8F2HoKZ5hmEW0aTMkuVYBHDE897kaWl5dymHksGf4mVHCqS251TWYTGW35qYHdw22A4toHqG94vTOWogryrsIhuQSUXyjCAnIO64AXG6e9hwr.JmpOeVpjWbKrP7yhqPIAstRWfkKWAlkN0D.s4JrlVX1PGn7DH3VHhVDkEOwX81FtVDWA4.B9F4OSfGMDgHJvcqMM8Ma5YttYqu4z.ebjLJILWfhm2fRWLQlaqpXq9U90WR63n5bHEi.MX3z3dTLbvho0h3OeKFYC5cJfBy9gF.kwUwCFcLeljE68Dt6HVv5D2XAmKuoZlINCPvVuzmfzCvVPv8AMddyx2FNbBfpH7R7mxB1lQClHYo0uQWz.1wXTlpfJI63HQKSnnSuQqDJUhxCSSoQO9LWejogAbl8PDeaCG.odXkDwoWwZzWNSztpcg_JBQ0YRQGIbvPYFhNdE_T70DTDeccFmR2fJBbzhslGqq7uRsnONwE_UyMZleaQpbWM.pJ.mUs6cWUl0CyO494Y4upBdGe3WRzH8Nbrzu7eP3eBQUKnf.Ijk5GJHfNY7fJx7EttRymdRnbXbWq9kYhpLw5cjsdGeqXnZeszHpwwtTE9lZTEuAcVzzaNVhLcwnYDAtrUumX0JdbGdKe2QHsCM25BFr1XQ6aiuy667TsK_apLBk5Mp4Kau3WzmIw60cCiVWPe6.Zf_qqWIdjb9uduPzGl7xO_dr3I0V7GdAk8WSz4VjjUhvk33e0xKeDZTxv3ZT_DnuoWZ7is5qSh4D_guzlXg_Jhg8bkElqOq5PqaJ.QT73TWcOdSgyMOQpnUOD6P10cDCOXeI9WgReKQAj_tP6uXudvS8fWDirRpuHKC5AtG1m_o5xV6od6fjQ6R6Ojl71SwwPu2CrezoxluMWR.JEM2cJbIwUW9w77XTeEjpNZv3Hc.zsYIiCa.60hLr9Lo6rQNXPaUPi_3Vo9FqfLVB.EzsPxAhLmoOTtxEz1R9Os9thH007bFY2PQAtA5aNt1kAps8edkp12Gn58DUbio5wx4wy6nOX4CQflQHVTh7zwm6uzHNTcCi6odxfn_H9oqll3myvLMU8a3Fw3vYRpaS6uW_dmA80U.A7k8HIEgMiSGvh5cHPLoKdPOG2s9FzDp0Jmm.6jw0KFNMpev_RMD4ia35.H0Nwo0BbwcqrUWPpphtuxJVv4.TLb2JQPmR_l2QWr_qsBpLQxVSO76HbA5WB_W3LgyHiVBcjHVLnLJaV93aBZnNXKHXNQAq1AB1xGjG4JWgGumzbMtaNVmkPIvxBH2zVuGAYzfYY5.D2gO9xT3K37U4w0TSnz0E1t12kYVk9ycKQpaLa1tRwKhM.pdKt_MJSeYDld1vVjbSiKtrDCYo8rewSkYsAQgA6PA96DfSJIg4lF1AmkXUyTv0ROMXn_VO5r_9zg.pLYBo1kAFCCmrIbFtnhyP4dZFaPQNX6pIvnXeFx8gXjxibmio3kwrdKkJS6m8L5Z8IqfpQYto5vU_6ekpN_y4nwDzmw9Z2PErnagTwLQKHeS3j.RnBANu5wxUrVWYPg3e5CO1NrrMf0mPCn0kZPFL0DT1kmlSAXHZSqoh3s3kqtgaVaugFnRLPPAACPrBhNl7_TR0vL_0qUb_xQn1e2UCcKRpM9FrUShX0iv8k9JJaIHRpyBA0MCMelXFPEeLxLm5g97nOXj.F1ZBQmSp5Ky9cum7XQHHtNvg_DuOsoFXvNU8aKqfUYQ7z8c0oXkUPO0ijSxSpFTH7cOW.uorkD5N7r2XqFPIvWerq93G2nghj6I2yNpQr38rL9f4h6h1kYhaIiUz4Y7WNfL7cZc987LMPEJbjv6VAutVrnRN69wFw8Q.HMZmXgP9GQ4RoWWI4WjS.gJrKzQl11QTpuVe50tepMZYVINWm2oJbgQHc0GPjoTlvlPkFfL1PBRi16JOHmu7fSvoxEezpAXtvTvtVQ16EyR_K6uWTM0FAsv0w9xazbeA8aUAYY4PBww_4.cvHroQZkBqecccq98_ZNqrMzHOPI5gI04.0YiSYjo.2XOIQBvtGPn4huj1INJGZma5hwe5voLCm6EiGJ.9R381x2q_0VEluDozBl.o7Rh2YatdzOIjg6wcf9DOWEta0ziu2Xkf0lbuo23tkAHDEYhWvpG3p.rNP0U8LkAJByKrb1rWcI3TaYRjUlQO_V_eZyY4gy1962D_z1.DpQcnaZkn.tK7jTnfPwzqdkY6DVRAfbCb4..6avFx8CTLotgkMari3eGra.aDCt632WHvqpk6QsP3SZ3.0OYLxZwgVRzmyGsXx1LDkNAPcmipLDOcbUoC7Y4Ot9dgEGNgqpHH5qY3HV6U170L_hsFsk83bu0F1yoGPPrG7kQDtA.n9aHgZP8e2Gvd5halC6cwnMW._Rvo38nlgxfa0F3FC2yT1ggy0TRTRZOd65tiWMrnkKVhDmkWJi.ZuLMjUd_Ou5IDKBxrnRm_bAcFHCx_Hlot0swiGG2PF8uYQDcKyVEGvl3f4ZS64e2pOzzykkYmM7A7Ur9f8UFQQ6nSKaJr2t_eKQutsk8segL6BhL0t3PZrHqxRcyJaz9RL7w0VJ90IL3tNlsYnW__iRd4Itg4XK.0dB.rEP1q5INeHuAmUYAvYK6TfkIrOPzc1Tnl.6ZWOUOmJ1kvk4XqeMHviBCKjSb5Iz56qORhUcTRRVGxOEaWJfXJFVj1.bvxMzGWl8OVCAGVzRupmLmicb5KdTDSroPtoH.0K8YHHPAnFQKU0Ro1JThjOonbHpH3RiN12lO9eU420a80ZqAyNuF9fvH9lYE.TIgMHX4OzTF8ysZd2YzkcHkkSFvTpZ2enJQkhLnvKIKr6fOjGrbmpSY7Yp6s6FJEkl6ZE7lyn81PbmK1CrqWk25qXxQ5E1WbRgZOd0Sf78aM3iDw1aWnIy2mC3Va.Dyd.MZ6UEhlvMTsTOAZNYmq0lkL8_suFJ6JZck",mdrd: "7aTPlReENiwUdVPQ1vzQf.eAs7QIp6vhoblykTRNI_A-1736140895-1.2.1.1-iGPfpTg82axbB3m72uTrHaYp1UoXRZPGi4vSls7jc09PLav87Un0Kuumz5EXYDoG2JSsHg2hHKV3dbxfMI.sMY84yikZ9L9iAScBsLi3QflRQfmPRvSChbE.p2BN.dmLEox6Dux99tRfO9u.bTYeVlC7oLxxSJq4pSvUISYDOXfgAtb0P8jCmdiZvlNV5z.ll.0M1c7uXPoa.o29mla6U1NSGlL5sYjG3xSmVQ62Ov_VZzNk9uvTPLhCJy6sJnJsgewwG3B0D94710tbvGTRvR6_iJHwrk0Ii_71knnKML1N4UPPes6mmWrRTHzk71zUGDUoMNojYYIsnjqE2a13ExkYYH.exWwcZ1Db90gEKz9AKoeVK38VCYmtoEh5WD08MFKS6DFExjobovw_7vjAW8hutWyvKOBKsQqi26jdM6miYYfqtGVQBnoxHEXj2HDdzIAyNkraiH2xqpcEY5fsW1nXHpVB3kq14SUd0kBx4BWtWL9NbwUZpWZA6yArjWHUcKGxbcTxpfXrn53WpCo46_92OZRO64mEqYbV_MdGB2CjTQzicSe7nRPtDM9Y_D7pUTSbEDKu64Ao6lV.OEk_kz4RZc57Y91QrQZDCcpsesuwd4p1_QWO3THE.3Ecz0Y2YRmq9itT4bp5NPLIVUhCo3VKTs1D4EG7qXhx3AD3WOEXr_AP_tBD791TQo06a7ZFgBpaIdVBPn5n7PjL9BMcklEAy6j4iA9O8x.C6Tv3JlCHR5CoZvgz8UQVlFL9XVX5vdNGcTscgsDVEUFyNoUduDhZg.s5pyvGozgbPTf_Ib11bvjqOxz1MTrvM7kUfUhLN0sZ.Jo8sQpKQX.CxRrma96sDegLVhzrAliq9_c5MZgTlcjcQzyb_CpIBumW05uEa1WR9FIdFh4j7dVKGTxTK0T9MLSEy7SDSfczR7SIp8KOf033bL5S6OCBzue38axu0HUQqE45M_ZhFLGDQO7OBYbER9k6dTD3.MFr4kRBJ1S9QMrCEuZZ7EsEF2hADlRxFxiVZBFZenVCN2aJw3PlwMtUcTb8jNqtvRD0k6wm__Cbyuj.NzgfHELjnqnTGabc4AArvUkM1.r0Y_JGmQSo4MLpMhR6XuW7fiCZ4wYxjgIcRuqbAeIB5_KY1BZJK9GelxixGlVRB.ovJ7mlzdJYLgQ_s8kfsMEUIQNardyD90Di8gu7bNVVMHgMnPNOpwlR3UG_hFBBKT_fNbAv.OFauG45Z6u3fwVH7H0yl2UrANcMGJZAh07Qfns1Sf3F9uDwgxJ4EGRSq5dny5eqx2l5dJZZRr82QioocyxisWGSHPUk_StQP6mjjxySSibfz6y5A_LSFhP5si4Nq4BBqjaFVmGwhEln9HPOuN0siClv3rPq3x5mqm6ZIWcLJjoJvZnwDukY7ysFoqdU0hjBApmfuluBiZdeqvpd9OMfJAL_t.2SxzAY45FFODkqDXD.vVpF8JPDlaVcwSIXSgZOLoqhAW2PktqkOKRpCP2DcFZm_33nGU6EywMIDBkwkhKSgWpbGgRIx4byCNk0p8NoyPEaaQbq.UvO2Gza91.jsfeuN5xZQPAfjNzCEIZgNKiy3GdoHy_ZEoq2JU4Xrj94JAgbWpc4wH8MB8VyF6db3NEz4jj9otwxE3liJRat2GjM4l1LWWB.rXGJYF13B166.fEySkMgMyfruZnyt6ttfx4LJl5_bdnu95iL3VsgGu06r3CULKj9dHxgpjQnJhX3knJcKZAfG.9h1mAinfvdwOvh9HZmKC6S4vzLCsQl2g3BW4n3a7tuHmNQeDfIPxdZYQveXn3ldd7KfAKJxpuDSY647Z3vaElf1iv0jjdSaA0hqMcqlB7a_m7SRJ8rjoZO.yeUI_cI1RDFQQH8zOjDiSN6BFE5C8epbwnJZ_5pAHrZIcdAMM6wwb7U6xXSt6_yz91zbAkR0tu.Ol237_VLNKl3ybR1A7IP73NNJ9zhCSOWTXEStHWehmrfaYIf7HarbqW5OQ"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/g/orchestrate/chl_page/v1?ray=8fd943f7bfa9021f';window._cf_chl_opt.cOgUHash = location.hash === '' && location.href.indexOf('#') !== -1 ? '#' : location.hash;window._cf_chl_opt.cOgUQuery = location.search === '' && location.href.slice(0, location.href.length - window._cf_chl_opt.cOgUHash.length).indexOf('?') !== -1 ? '?' : location.search;if (window.history && window.history.replaceState) {var ogU = location.pathname + window._cf_chl_opt.cOgUQuery + window._cf_chl_opt.cOgUHash;history.replaceState(null, null, "\/doi\/abs\/10.1111\/gcb.15913?__cf_chl_rt_tk=vIEGH1hh8sID48KObbzsNggu0n2iA_nbvt72Y_p.bHk-1736140895-1.0.1.1-2JAYcK59dTpFbzC1mwZe35ArUNa7OBY1gZaNtF2oSSs" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>